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THE AUSTRALIAN 


ntomologist 


published by 
THE ENTOMOLOGICAL SOCIETY OF QUEENSLAND 





Volume 25, Part 1, 5 June 1998 
Price: $5.00 per part 





Published by: THE ENTOMOLOGICAL SOCIETY OF QUEENSLAND 
ISSN 1320 6133 











THE AUSTRALIAN ENTOMOLOGIST 


The Australian Entomologist (formerly Australian Entomological Magazine) is & 
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Cover: Anthrax maculata (Diptera: Bombyliidae) described by Macquart in 1846 
has been collected commonly throughout eastern Australia, across northern 
Australia and south-western W.A. Specimens have been collected flying around 
burnt trees and mud wasp nests. Females are a common sight in suburban 
Brisbane, patrolling brick walls searching for mud wasp nests. Illustration by 
Chris Lambkin, Department of Entomology, University of Queensland. 


Australian Entomologist, 1998, 25 (1): 1-6 1 


OBSERVATIONS ON THE BIOLOGY OF ARHOPALA WILDEI 

MISKIN (LEPIDOPTERA: LYCAENIDAE) AND ITS HOST ANT 

POLYRHACHIS QUEENSLANDICA EMERY (HYMENOPTERA: 
FORMICIDAE) 


Rod Eastwood’ and Allan J. King’ 


150 Broadwater Terrace, Redland Bay, Qld 4165 
?PO Box 1302, GPO Townsville, Qld 4810 


Abstract 

Observations on the biology of Arhopala wildei Miskin and Polyrhachis queenslandica Emery 
are recorded, as well as details of their interactions. Ant assisted cuticle removal during ecdysis 
of A. wildei larvae is recorded for the first time, as well as pupal stridulation and pupal 
oscillation (vibration) in A. wildei. 

Introduction 

The myrmecophagous early stages of the lycaenid butterfly, Arhopala wildei 
Miskin, have been described recently by King and Ring (1996). The 
butterfly deposits its eggs near the nest entrance of the arboreal ant 
Polyrhachis queenslandica Emery and the first instar larvae are carried into 
the nest by the ants. The larvae then remain in the nest where they feed on 
the ants’ brood. Additional behavioural observations reported in this paper 
were made independently by the present authors at the same location near 
Waughs Pocket, north of Innisfail and at other sites in northern Queensland. 


Observations 


Adult butterfly behaviour 

Males of A. wildei are active in early morning sunshine (c >0600h EST), 
often flying close to the ground in clearings whilst “dogfighting” with other 
males. During the early part of the day males often search the foliage of vine 
forest trees, especially near P. queenslandica nests, possibly in search of 
freshly emerged females. They later retreat into the foliage and are seen only 
occasionally. Pairs in copula were observed on exposed foliage in mid- 
morning (c 0900h EST), adjacent to areas of male activity. When disturbed, 
mating pairs drift to a lower perch, seemingly unable to sustain flight. The 
mating ritual was not observed. 


Females are seen most commonly investigating the foliage between mid- 
morning (0900h) and mid-afternoon (1500h) and have been observed 
depositing eggs between these times. Whilst searching for egg laying sites, 
females adopt a slow flight pattern with much fluttering in and around the 
foliage. On occasion, they will fly quickly out of the foliage with a short 
burst of fast circling flight — apparently after being disturbed. Some older 
females remain in the vicinity of P. queenslandica nests for extended periods 
(3-4 hrs), occasionally flying off and returning to rest nearby. 


Adult butterflies were not seen to visit any flowers in the study area. 
However, they were seen in the company of other Arhopala Boisduval 


2 Australian Entomologist, 1998, 25 (1) 


species, A. micale amphis Waterhouse and A. madytus Fruhstorfer and other 
insects (wasps and ants), feeding on exudates at the leaf petiole junction of 
the large-leafed vine Merremia peltata (L.) Merr. (Convolvulaceae). 


The ant nest 

Nests of P. queenslandica ranged in size from very small (c 1 cm’), 
containing just a Queen, sometimes with a few early stages, up to large nests 
(>100 cm’) with more than 100 adult ants and a similar number of immature 
stages. Most nests were around 30 cm’ and contained 40-50 mature ants. A 
typical P. queenslandica nest is constructed of two, or sometimes three 
overlapping leaves sealed along the sides with what appears to be masticated 
bark. The inside of the nest has no compartments and all surfaces are 
covered by a thin layer of light brown silk. A dead pupa of A. wildei in one 
P. queenslandica nest was likewise webbed over. Two or three tube-like 
access holes of 3 mm diameter are built into the webbing at opposite ends of 
the nest. During the day, guard ants can be seen in these access holes. 


P. queenslandica also nest opportunistically. One group nested in a cavity 
under a plastic table using nesting material removed from a damaged nest. In 
the wild they have been found nesting in an abandoned wasp nest (S. 
Robson, pers. comm.). 


Nest sites for P. queenslandica varied in height from 30 cm above ground to 
high in the intermediate canopy in protected positions. Most of the nests 
were situated 3-5 m above the ground in the overhanging lower canopy of a 
forest boundary. This part of the canopy is protected from strong winds and 
is also where most of the A. wildei females were observed to fly. Ant nests 
may be up to 30 m above the ground as some A. wildei females were seen 
investigating foliage at this level. P. queenslandica nests were found on both 
windward and leeward sides of forest boundaries, in foliage with and without 
green tree ants Oecophylla smaragdina (Fabricius) and other Polyrhachis F. 
Smith species. 


P. queenslandica nests are very clean inside. Remains of ant eggs, larvae 
and pupae after consumption by A. wildei larvae are ejected from the nest by 
the ants. Dead ants and A. wildei larval frass also are ejected. However, 
some nests contained a number of fluffy, white, papery objects of various 
sizes which proved to be the masticated exuviae of A.wildei larvae. No 
empty A. wildei pupal cases were found in an active nest and it would appear 
that the ants destroy or eject empty butterfly pupal shells. 


Ant behaviour 

When disturbed, P. queenslandica adopt a defensive threatening posture. 
They curl their gaster forward under the mesosoma, striking it repeatedly on 
the substrate for one to two seconds, producing an audible rattling or 
drumming sound. In daylight hours, the ants are reluctant to leave the nest, 
even when disturbed. If the nest is breached, workers will form a line, face 


Australian Entomologist, 1998, 25 (1) 3 


the exposed breach in the defensive posture and spray fluid from the gaster at 
any intruder. Observations made on a number of disturbed nests showed that 
nests suffering minor damage were quickly repaired the same night, while the 
majority of those that had sustained major damage were found abandoned the 
next day. 


When two ant nests taken from about 50 m apart were placed in the same 
container, these nests merged into one with the expulsion of one queen and 
three worker attendants. The exiled queen was later killed by two other 
workers and the three attendants returned to the new single nest. On other 
occasions, ant brood from one nest was placed in a container housing another 
ant nest taken from a different location. The introduced ant brood was 
picked up by the other ants and placed inside the nest. P. queenslandica 
from different nests were not aggressive to one another as is the case with 
many other species of ants (Hdlldobler and Wilson 1990). A. wildei larvae 
transferred from one nest to another were similarly accepted without 
hesitation. 


Butterfly oviposition 

A. wildei eggs are laid singly or in small groups of two to four on the ant nest 
material, usually near the tubular access holes. The host ants were not seen 
to attack females laying eggs. A small number of P. queenslandica nests, 
with and without A. wildei eggs attached, were found to be abandoned and 
one abandoned ant nest was found to contain an empty pupal shell of A. 
wildei. Nests without eggs attached were occasionally found to contain one 
or more semi-mature A. wildei larvae. More often there were fewer butterfly 
larvae in the ants’ nest than hatched eggs on the outside. The majority of 
nests with butterfly eggs were around 20-30 cm’ and had 3-5 eggs attached. 
Some of the large-leafed evergreen trees in which larger nests were found, 
hold their foliage for at least two years and it is possible that these nests have 
supported continuous generations of A. wildei over that period of time. Only 
one intact A. wildei egg was found with a hole cut in the side, indicating that 
egg parasitoids may be present. 


A. wildei larvae in the ant nest 

In similar fashion to another myrmecophage, Acrodipsas illidgei Waterhouse 
& Lyell (Samson 1989), freshly emerged first instar A. wildei larvae are 
carried into the nest by the ants and therein remain very difficult to detect. 
Eggshells are neither eaten by the freshly emerged larvae nor removed by the 
ants and oophagy was not observed. A. wildei larvae of less than 2 mm in 
length were seen clinging to the ants’ silken pupal cases or attached to ant 
eggs in the brood batch. Ant eggs and larvae appear to be coated with a thin, 
viscid layer. This substance enables the ant eggs and larvae to adhere to each 
other and to the nest wall, and facilitates the movement of batches of brood 
by individual ants. Small A. wildei larvae also adhere to the ant brood and 
are likewise moved by the ants. In the event of a major disturbance the ants 


4 Australian Entomologist, 1998, 25 (1) 


pick up the ant brood, with lycaenid larva attached, and retreat to the darker 
corners of the nest. Ants will often pick up A. wildei larvae instead of their 
own brood. 


The host ants do not show any aggressive behaviour towards any instar 
larvae of A. wildei. A. wildei larvae have been observed consuming P. 
queenslandica eggs and larvae and there is circumstantial evidence that they 
also eat ant pupae. Trophallactic feeding was not observed between the 
butterfly larvae and adult ants. However, it may occur since the. ants. seem to 
treat A. wildei larvae, in all other respects, like their own. Butterfly larvae 
are regularly attended by the ants with particular attention being paid to the 
Newcomer’s organ (NO). Exudates from this organ collect in the concave 
anal depression of the A. wildei larva and often, when approached by an ant, 
the larva will raise its posterior end to a vertical position, facilitating direct 
access to the NO by the ant. Similar behaviour has been observed in 
Acrodipsas illidgei by Samson (1989). The queen ant also attends the NO of 
A. wildei. 


A. wildei larval ecdysis 

Two attendant P. queenslandica were observed to carefully remove the 
moulting cuticle from a second instar A. wildei larva. The action was 
performed at first with both ants pulling and then with one ant holding the 
larva in its mandibles, while the other ant slowly peeled back the loose 
exuvium. This was followed by a lengthy session of meticulous grooming 
with the larva being turned over several times, apparently under duress. The 
exuvium was then picked up by one of the attendant ants and thoroughly 
masticated before being dropped. It was not ejected from the nest. This 
entire process was performed by the same two ants without intervention from 
others nearby. 


Pupal stridulation and oscillation (vibration) 

A. wildei pupae stridulate but, unlike other lycaenids that stridulate with 
intermittent bursts of “ticks” or “burrs”, they emit a prolonged “burrr” that 
may last for two to three seconds. This was particularly noticeable after a 
pupa had been removed from and subsequently reintroduced to an ant nest. 
The pupa made no noise until it was “attended” by the ants. This 
phenomenon of pupal stridulation being immediately induced by contact with 
an attendant ant also occurs in other species of lycaenids, e.g. Jalmenus 
evagoras (Donovan) (RE, unpublished observation). 


A. wildei pupa were also observed to oscillate. This oscillation consisted of a 
rapid dorso-ventral movement of the anterior end of the pupa. It was not 
determined if the pupal oscillation coincided with the sound production but 
they appeared independent. Interestingly; the frequency of the pupal 
oscillations also appeared to be the same as the frequency with which the 
host ants tapped on the substrate when alarmed. 


Australian Entomologist, 1998, 25 (1) 5 


Discussion 

Adults of A. wildei dramatically vary in size with wingspans of males and 
females ranging from 25-41 mm. The smallest adults may result from semi- 
mature larvae that had exhausted their available food supply or were left 
behind when a nest was abandoned by the ants, forcing the lycaenid larvae to 
pupate early. There is no evidence to indicate that larger A. wildei larvae 
migrate to new ant nests as is the case with another arboreal 
myrmecophagous lycaenid, Liphyra brassolis Westwood (Dodd 1902, J. 
Young pers. comm.). Smaller A. wildei larvae may be carried by P. 
queenslandica to a new site should the ant nest be destroyed or abandoned. 
However, larger larvae are very vulnerable and desiccate quickly outside the 
ant nest and may not be able to survive an extended journey. 


Another measure that may be used by the butterfly to ensure an adequate 
food supply is cannibalism. In smaller ant nests or under adverse conditions 
A. wildei larvae seem to be able to regulate their numbers, and hence the 
available food supply, by this method. This hypothesis is supported by 
circumstantial evidence, including the fact that some smaller P. 
queenslandica ant nests were found to contain only one lycaenid larva yet 
had 2 or 3 recently eclosed eggs attached. 


A very similar behaviour to the ant-assisted cuticle removal was recorded by 
Brewster (1913), where two Polyrhachis ammon (Fabricius) ants assisted the 
eclosion of a winged ant from its pupa. After being removed from the pupa 
the two ants continued to assist the imago “...one holding while the other 
pulled the wings clear of their sticky covering.” Traniello (1982) and 
Hölldobler & Wilson (1990) also describe the care that ants bestow on their 
eggs, larvae and pupae including that of assisting the larval molt by licking 
the ecdysial skin free. Eclosion of an A. wildei adult in the presence of ants 
was not observed. 


Pupal noise, produced by hammering on the substrate, has been recorded in 
six species of lycaenids (Downey 1966) and he suggested that the function 
was to frighten away small predators. However, DeVries (1990) suggested 
that lycaenid pupal noises mimic vibrations used by the ants for 
communication. It is possible that the pupal oscillation of A. wildei is a 
specialised form of communicative stridulation or alarm response similar to 
that of the P. queenslandica ants. l 


Pierce et al. (1987) deduced that significant extra numbers of new workers of 
Iridomyrmex anceps (Roger) can be produced by ants that are tending 
lycaenids and Nash (1989) was able to provide direct evidence that ant 
colonies exhibited significantly higher growth rates when they were able to 
tend lycaenid larvae. It would be reasonable to assume that larvae of A. 
wildei can similarly stimulate P. queenslandica. In addition, the NO of late 
instar larvae of A. wildei is proportionally larger than those observed on other 
myrmecophilous lycaenids, such as J. evagoras (Kitching 1983). If it is the 


6 Australian Entomologist, 1998, 25 (1) 


NO that produces the compounds that stimulate fecundity in the attendant 
ants, then this organ would be a very valuable asset to the A. wildei larva and 
may account for its increased size and prominence. Of course it is entirely 
possible that the exudates from the A. wildei organs are simply to ‘bribe’ the 
ants to prevent predation (Malicky 1970), while the larva consume the ants’ 
brood. Whatever the purpose of the exudates, they are evidently very 
attractive to the ants. The exudates are also persistent, apparently remaining 
on the moulted A. wildei exuviae in sufficient amounts so that the ants are 
reluctant to eject these “foreign” objects from the nest. 


Acknowledgments 

We gratefully acknowledge Lance and Lyn Vievers for permission to study 
and film A. wildei on their property at Waughs Pocket. Thanks also to Ann 
Fraser for helpful comments on an earlier draft of this paper and to Rupert 
Barrington and Alan Hayward (both BBC London). 


References 

BREWSTER, M.N. 1913. Observations on ants. Australian Naturalist 2: 178-179. 

DEVRIES, P.J. 1990. Enhancement of symbioses between butterfly caterpillars and ants by 
vibrational communication. Science 248: 1104-1106. 

DODD, F.P. 1902. Contributions to the life-history of Liphyra brassolis, Westw. The 
Entomologist 35: 184-188. 

DOWNEY, J.C. 1966. Sound production in pupae of Lycaenidae. Journal of the 
Lepidopterists’ Society 20(3): 129-155. 

HOLLDOBLER, B and WILSON, E.O. 1990. The ants. Belknap Press of Harvard University 
Press: Cambridge; 732pp. 

KING, A.J. and RING, L.R. 1996. The life history of Arhopala wildei wildei Miskin 
(Lycaenidae). Australian Entomologist 23(4): 117-120. 

KITCHING, R. 1983. Myrmecophilous organs of the larva and pupa of the lycaenid butterfly 
Jalmenus evagoras (Donovan). Journal of Natural History 17: 471-481. 

MALICKY, H. 1970. New aspects on the association between lycaenid larvae (Lycaenidae) 
and ants (Formicidae, Hymenoptera). Journal of the Lepidopterists’ Society 24: 190-202. 
NASH, D. 1989. Cost-benefit analysis of a mutualism between lycaenid butterflies and ants. 
Dissertation. University of Oxford, Oxford, UK. 

PIERCE, N.E., KITCHING, R.L., BUCKLEY, R.C., TAYLOR, M.F.J. and BENBOW, K.F. 
1987. The costs and benefits of cooperation between the Australian lycaenid butterfly, 
Jalmenus evagoras, and its attendant ants. Behavioral Ecology and Sociobiology 21: 237-248. 
SAMSON, P. 1989. Morphology and biology of Acrodipsas illidgei (Waterhouse and Lyell), a 
myrmecophagous lycaenid (Lepidoptera: Lycaenidae: Theclinae). Journal of the Australian 
Entomological Society 28: 161-168. 

TRANIELLO, J.F.A. 1982. Population structure and social organization in the primitive ant 
Amblyopone pallipes (Hymenoptera: Formicidae). Psyche 89(1-2): 65-80. 


Australian Entomologist, 1998, 25 (1): 7-12 7 


A NEW SPECIES OF TRAPEZITES HUBNER (LEPIDOPTERA: 
HESPERIIDAE) FROM WESTERN AUSTRALIA 


Andrew A.E. Williams‘, Matthew R. Williams’ and R.W. Hay’ 


'Department of Conservation and Land Management, W.A. Wildlife Research Centre 
PO Box 51, Wanneroo, WA 6065 


*Department of Conservation and Land Management, 50 Hayman Road, Como, WA 6152 
’8 Klem Avenue, Manning, WA 6152 


Abstract 

Trapezites atkinsi sp. nov. is described from south-western Western Australia and, on present 
knowledge, is-limited to a small area of coastal heathland at Windy Harbour. The pupal stage 
is described and illustrated. The larval foodplant is Acanthocarpus preissii Lehm. 
(Dasypogonaceae). 

Introduction 

In reviewing the Western Australian species of Trapezites Hiibner, Mayo and 
Atkins (1992) drew attention to a form from Windy Harbour, which they 
tentatively assigned to the T. sciron complex. Specimens of this skipper 
were first collected by David Yeates on 11 November 1989; subsequently 
further specimens were taken in 1989, 1990 and 1995. Examination of this 
additional material confirmed earlier suspicions that the original specimens 
represented a previously unrecognised species of Trapezites. T. atkinsi sp. 
nov. is closely allied to three other Western Australian species of Trapezites: 
T. sciron Waterhouse & Lyell, T. argenteoornatus (Hewitson) and T. 
waterhousei Mayo & Atkins. 


Abbreviations 

The following abbreviations refer to institutional and private collections: 
AA = Andrew Atkins Collection, Newcastle; RWH = R. W. Hay Collection, 
Perth; HHB = H. H. Bollam Collection, Chittering; WADA = Western 
Australian Department of Agriculture Collection, Perth; CALM = 
Department of Conservation and Land Management Collection, Perth; 
WAM = Western Australian Museum, Perth. 


Trapezites atkinsi sp. nov. 
(Figs 1-9) 


Types. WESTERN AUSTRALIA: Holotype d', Windy Harbour, 29.x.1995, 
low coastal heath above cliffs with Acanthocarpus, 34°50’13”S 
116°00’49”E., A.A.E. Williams, Reg. No. WAM 1998/0013 (in WAM). 
Paratypes (11 0”, 8 2): 1 9, Windy Harbour, 14.xi.1995, ex pupa in shelter on 
Acanthocarpus preissii, 34°50’13”S 116°00’49”E., A.A.E. Williams, Reg. 
No. WAM 1998/0014 (in WAM); 1 2, Windy Harbour, 8.xi.1995, ex pupa in 
shelter on Acanthocarpus preissii, 34°50°13”S 116°00’49”E., A.A.E. 
Williams; 1 0’, Windy Harbour, 8.xi.1990, M.R. Williams; 1 0%, Windy 
Harbour, 29.x.1995, M.R. Williams; 2 9, Windy Hbr., 34°50’13”S 
116°00’49”E., ex pupa, 2 & 7.x1.1995, M.R. Williams (all in CALM); 1 ©’, 


8 Australian Entomologist, 1998, 25 (1) 


Windy Harbour, 26.xi.89, R.H.; 1 d, Windy Harbour, 8.xi.1990, M.R. 
Williams (RWH); 3 0’, Windy Harbour, Pt D’Entrecasteaux, 19.xi.1989, D. 
Yeates, each with an additional label, Agriculture (Dept) Western Australia, 
with an identifying database number 41347, 41348 or 41349; 1 9, same data 
but number 41350, with genitalia in attached vial; 2 0’, Windy Harbour, 
24.xi.1989, H. Bollam, both with additional labels as above but numbered 
41345 and 41346, both with genitalia in attached vials (all in WADA); 1 CO’, 
Windy Harbour form, xi.1989, H. Bollam; 1 0’, Windy Harbour, 8.xi.1990, 
M.R. Williams; 1 9, Windy Harbour, 29.x.1995, reared ex pupa, A. Atkins, 1 
9, Windy Harbour, 1.xi.1995, reared ex pupa, A. Atkins; 1 9, Windy 
Harbour, 6.xi.1995, reared ex pupa, A. Atkins (all in AA). 


Male (Fig. 4). Head dark brown with hair-tufts of light brown, beneath 
greyish-cream; antennal shaft dark brown above, segmented with yellowish 
scales, greyish-cream beneath with darker segments, club dark brown to 
black, nudum 14-15 segmented black; labial palpus black above, cream 
beneath, third segment short and black. Thorax above black with brown hair 
scales, posterior with long yellowish-grey hair-scales; legs pale brown, hind 
leg often with two pairs of spurs. Abdomen brown, with distal edges of 
segments pale yellowish-brown, posterior hair-tuft pale brown, beneath 
yellowish-grey. Forewing length 14 mm, with costa almost straight, apex 
pointed, termen almost straight; above centrally mid-brown to dark brown 
with pale greyish yellow hairs on termen, base and central (median) area with 
yellowish grey hairs, three to four subapical pale yellow spots in area 
between R3, R4, R5 and M1 diagonally placed across wing, a yellowish cell 
spot and two yellowish-orange spots, one between M3 and CuAl and a 
slightly larger spot between CuAl and CuA2, two confluent yellowish- 
orange spots forming an arc or crescent in median area between CuA2 and 
1A+2A, sometimes reduced or nearly absent; cilia pale grey-brown; beneath, 
costa, apex, inner margin to tornus and small area above median spots 
yellowish-grey, distad of which are two dark spots, median area dark brown 
to black, all spots as above but slightly paler; cilia cream with slightly darker 
chequering. Hindwing slightly rounded, above dark brown, yellowish hairs 
arising from base to median area and extending along inner margin to 
subtornal area, a large central yellow-orange patch of scales; cilia pale 
greyish-yellow; underside pale yellowish-grey, basal area greyish-brown to 
grey, central patch as above but pale yellow, six subterminal and three 


Figs 1-9. Pupal stage and adults of Trapezites atkinsi sp. nov. (1) frons of pupa; (2) 
lateral and dorsal view of pupa; (3) pupal setae; (4) adult male (left upperside, right 
underside); (5) adult female (left upperside, right underside); (6) female genitalia 
(dissected from specimen #41350 in WADA collection); (7) male genitalia including 
outside left valva (dissected from specimen labelled Windy Harbour form, xi.1989, 
H. Bollam, in AA collection); (8) uncus (ventral); (9) posterior section of outside 
right valva. Scale bars: Figs 1-2 = 5 mm; Fig 3 = 0.5 mm; Figs 4-5 = 10 mm; Fig 6 = 
1 mm; Figs 7-9 = 2 mm. 


Australian Entomologist, 1998, 25 (1) 





10 Australian Entomologist, 1998, 25 (1) 


median black spots centred with silvery-white scales (the largest subterminal 
spot being near tornus); cilia cream with slightly darker chequering. 


Genitalia (Figs 7-9). Uncus/tegumen broad with uncus (Fig. 8) tapered, 
rounded and blunt, slightly toothed at ‘T’-shaped tip, gnathos broadly 
rounded and moderately toothed distally, dorso-lateral flanges short, simple 
and distally rounded, directed slightly posterior; saccus short, beak-shaped, 
connected to a simple, broadly curved vinculum; left valva broad, toothed 
and sclerotized posterio-dorsally, harpe, broad, slightly dentate and short 
sacculus directed dorsally, ampulla rounded and blunt, right valva slightly 
more dentate and toothed; aedeagus short and broad, posterior tip (postzone) 
expanded, semi-cupped shape; juxta simple and saddle-shaped. 


Female (Fig. 5). Similar to male, but forewing and hindwing with apex and 
termen more rounded, spots on forewing larger, with the two confluent spots 
forming a crescent between CuA2 and 1A+2A prominent. Spots on both 
wing surfaces larger and more prominent. Forewing length 15 mm. 


Genitalia (Fig. 6). Papilla analis broad and triangular, apophysis long, 
straight and slender; sterigma plates with lamella postvaginalis elliptical, 
long, thin and steeply ‘V’-shaped with deeply rounded central groove, 
lamella antevaginalis simple, broadly ‘U’-shaped and slightly sclerotized; 
caudal chamber moderately broad and short; ductus bursae short and 
spreading; corpus bursae broadly ovoid with creased and slightly sclerotized 
base and a short, spherical accessory pouch attached by a short, narrow neck. 


Distrbution 
Trapezites atkinsi is known only from a small area of coastal heathland at 
Windy Harbour, Point D’Entrecasteaux, in south-western Western Australia. 


Etymology 
The species is named in honour of Mr Andrew Atkins, in recognition of his 
contributions to Australian entomology over many years and particularly his 
contribution to the study of Hesperiidae. 


Variation 

There is slight variation in the size of the median spot along the inner margin 
of the forewing; generally this is distinctively crescent-shaped. In some 
specimens the underside of both wings is more evenly pale yellowish-brown. 
There is also slight variation in the size of the spots on the underside of the 
hindwing. 

Life History 

Foodplant. Acanthocarpus preissii Lehm. (Dasypogonaceae). 


Pupa (Fig. 2). Length 18 mm; greyish-brown covered with a prominent 
mottled dark brown maculation and branched setae (Fig. 3); pupal cap 
(operculum) (Fig. 1) rough, sclerotized and subelliptical to quadrate, lightly 


Australian Entomologist, 1998, 25 (1) 1] 


covered with branched setae and mottled maculation, upper area extended to 
quadriform ‘turret’, lower area strongly dentate; cremaster short, tapered and 
decurved with rounded tip. 


Discussion 

T. atkinsi resembles all other Western Australian species of Trapezites in 
wing-shape, general pattern and colour of both upperside and underside. It is 
most similar to T. argenteoornatus in having prominent yellowish-orange 
maculation, especially that of the upperside of the hindwing. However, T. 
atkinsi is somewhat larger, the wings lack prominent chequered cilia, and the 
silver spots on the underside of the hindwing are smaller and on a yellowish- 
grey ground. 


The broadly rounded valvae tips, ampulla, broadly rounded gnathos and 
dorso-lateral processes of the uncus in the genitalia distinguish males of this 
species from other Western Australian Trapezites species (Mayo and Atkins 
1992, Andrew Atkins, pers. comm.). Female genitalia of T. atkinsi are 
differentiated by the very long, flat ovoid extensions to the lamella post 
vaginalis plate and a broad ‘U’-shaped lamella antevaginalis plate. In profile 
the genitalia of both sexes appear closer to T. sciron eremicola Burns than to 
T. argenteoornatus. 


The pupa of T. atkinsi is strongly mottled and resembles that of T. 
argenteoornatus, but the operculum is higher and dentate at the base and has 
a distinctive raised dorsal area. Larval and pupal shelters are similar to those 
of T. argenteoornatus. Both species are recorded on the foodplant A. preissii 
but T. argenteoornatus has not been recorded south of Bunbury. 


Larvae and pupae have been found in shelters on the foodplant or on nearby 
vegetation. The foodplant grows along coastal dunes and on the top of 
limestone headlands. Adults fly in areas around the foodplants during late 
October and November. They “patrol” in sunshine and often settle on 
sheltered sandy patches. 


The discovery of T. atkinsi brings the number of Trapezites species found in 
south-west Western Australia to four, with three of these being restricted to 
this area. The phylogeny of this interesting group would no doubt reward 
further study, as they all are very closely related. Three (T. waterhousei, T. 
atkinsi and T. argenteoornatus) are allopatric, whereas the fourth (T. sciron) 
may be sympatric or parapatric. Closer study of this group may shed further 
light on the complex variation observed in T. sciron and T. argenteoornatus 
by Mayo and Atkins (1992). 


Conservation 

T. atkinsi is found within D’Entrecasteaux National Park, but its apparently 
restricted distribution places this skipper in a vulnerable category. Our 
searches south-east of Windy Harbour and in seemingly identical habitats 


12 Australian Entomologist, 1998, 25 (1) 


between Yallingup and Cape Naturaliste have failed to detect further 
populations of the skipper. 


Common name 
In order to ensure consistency with proposed changes to the common names 
of Australian butterflies, we suggest the common name “Heath Ochre” for T. 


atkinsi. 


Acknowledgments 

We are grateful to David Yeates for information and discussion, to Hugh 
Bollam for records and material, and to John van Schagen for lending 
material held in the Western Australian Department of Agriculture collection. 
We thank Andrew Atkins for genitalia dissections and for providing the 
illustrations. Valuable comments on the manuscript were received from 
Michael Braby. 


Reference 

MAYO, R. and ATKINS, A. 1992. Anisyntoides Waterhouse (Lepidoptera: Hesperiidae): a 
synonym of Trapezites Hiibner, with description of a new species from Western Australia. 
Australian Entomological Magazine 19: 81-88. 


Australian Entomologist, 1998, 25 (1): 13-22 13 


NEW LARVAL FOOD PLANTS FOR AUSTRALIAN HAWK MOTHS 
(LEPIDOPTERA: SPHINGIDAE) 


M.S. MOULDS 
Entomology Department, Australian Museum, 6-8 College St, Sydney, NSW 2000 


Abstract 

New food plants are listed for 31 species of Australian hawk moths. Fifty-five of these food 
plant records are of native plant species and 17 are exotics. Three previously published food 
plant records for Cephonodes kingii (W.S. Macleay) and one for Psilogramma menephron 
(Cramer) are disputed, together with one for Hopliocnema brachycera (Lower) that originates 
from a labelled museum specimen. A brief overview of the diversity of Australian hawk moth 
food plants is given. 


Introduction 

In previous papers (Moulds 1981, 1984) I summarised larval food plants for 
Australian hawk moths affecting garden ornamentals and commercial crops. 
However, many Australian hawk moth species have larvae that feed on 
neither ornamentals nor crops and this paper lists known food plants for a 
number of these in addition to unrecorded food plants for species treated 
previously. Food plants listed by Common (1990) are not repeated. 


The following abbreviations are used when listing names of observers with 
several records: AJG, Alan Graham; GS, G. Sankowsky; JO, J. Olive; MSM, 
M. S. Moulds. 


Nomenclature for sphingid species follows that of Moulds (1996). Plant 
names follow Henderson (1997), otherwise Brock (1988) or Bailey et al. 
(1976). Exotic plant species are marked by an asterisk (*). 


Food plant identifications originating from H. Beste, AJG, GS, D. Lane, 
MSM, JO, C. Pratt, P. Valentine and A. Walford-Huggins were provided by 
Tony Irvine, CSIRO, Forest Research Station, Atherton or Garry Sankowsky, 
Yuruga Nursery, Walkamin. Moth identifications were obtained by rearing 
larvae to adults; doubtful adult identifications were confirmed by the author. 


New records 


Acosmeryx miskini (Murray) 
VITACEAE 
Cayratia clematidea (F. Muell.) Domin "wild grape" [GS, Tolga, Qld; A. 
Hiller, Mt Glorious, Qld] 


Agrius godarti W.S. Macleay 
CONVOLVULACEAE 
*Ipomoea batatas (L.) Lam. "sweet potato" [MSM - Larvae were not found 
naturally on this plant but several reared from eggs successfully developed to 
adults. ] 


14 Australian Entomologist, 1998, 25 (1) 


Cephonodes hylas (Linnaeus) 
Add to the records listed by Moulds (1984) the following: 


RUBIACEAE 
Pavetta granitica Bremek. [D. Lane and MSM, Dimbulah; GS, Tolga, Qld] 


Psychotria sp. "wild coffee" [Anne Garrett, Rockhampton district, Qld] 


Cephonodes kingii (W.S. Macleay) 
Add to the records listed by Moulds (1984) and Common (1990) the 
following: 


RUBIACEAE 
Gardenia ochreata F. Muell. "scented gardenia bush" [GS, Chillagoe and 
Georgetown, Qld] 


Gardenia ovularis F. M. Bailey [GS, Tolga, Qld] 
Tarenna sp. [GS, Tolga, Qld] 


NOTE: Jones and Elliot (1995) list Cissus, Grevillea and Oreocallis as food 
plants. Oreocallis was previously known as Embothrium and now as 
Alloxylon. All stem from old inaccurate records and should be disregarded 
(see Moulds 1984: 60). 


Cephonodes picus (Cramer) 
Add to the records listed by Moulds (1984) and Common (1990) the 
following: 


RUBIACEAE 
Aidia racemosa (Cav.) Tirveng. [Anne Garrett, Rockhampton district, Qld] 


Coenotes eremophilae (T. P. Lucas) 
Add to records listed by Moulds (1984) and Common (1990) the following: 


ACANTHACEAE 

*Barleria cristata L. "Philippine violet" [E. A. Henty, Kununurra, WA] 
RUBIACEAE 

*Mussaenda sp. [E. A. Henty, Kununurra, WA] 

VERBENACEAE 


Stachytarpheta urticifolia (Salisb.) Sims "ratstail" "dark blue snakeweed" [E. 
A. Henty, Kununurra, WA] 


Vitex glabrata R. Br. [Cliff Meyer, Kununurra, WA] 


Coequosa triangularis (Donovan) 
Add to the records listed by Moulds (1981) the following: 


PROTEACEAE 
Grevillea asplenifolia x caleyi 'ivanhoe' [N. Marks, Pennant Hills, NSW] 


Australian Entomologist, 1998, 25 (1) 15 


Hakea spp. [McMaugh (1985, 1986)] 


Persoonia levis (Cav.) Domin [Bruce White, Doyalson, NSW - Rose (1975) 
fed P. levis to newly hatched larvae which died. The discovery of larvae 
feeding naturally on this plant by White confirms it is a larval food plant.] 


Daphnis hypothous (Cramer) 
APOCYNACEAE 
Alstonia actinophylla (A. Cunn.) K. Schum. "milkwood" [G. Brown and 
MSM, Darwin, NT] 


Alstonia constricta F. Muell. "bitterbark" [GS, Tolga, Qld] 

Alstonia muelleriana Domin "hard milkwood" [MSM, Julatten, Qld] 
Alstonia scholaris (L.) R. Br. "milky bean", "milky pine" [MSM, Julatten, 
Qld] 


RUBIACEAE 

*Anthocephalus chinensis (Lam.) A. Rich ex Walp. [Gary Fitt, Darwin, NT - 
a tree grown experimentally for commercial use near Darwin; larvae 
completely defoliated some plants.] 


Nauclea orientalis (L.) L. "Leichhardt tree" [Gary Fitt, Kununurra, WA; P. 
Valentine, Townsville, Qld] 

Daphnis placida (Walker) 
Add to the records listed by Common (1990) the following: 


ALANGIACEAE 
Alangium villosum subsp. polyosmoides (F. Muell.) Bloemb. [J. Stockard, 
Wingham Brush, NSW] 


Alangium villosum subsp. tomentosum (F. Muell.) Bloemb. [GS, Burnett R. 
and Wallaville, Qld] 


APOCYNACEAE 
Alstonia actinophylla (A. Cunn. ) K. Schum. [C. Pratt, MSM, Cooktown, Qld; 
G. Brown, Darwin, NT] 


Alstonia muelleriana Domin "hard milkwood" [MSM, Julatten, Qld] 
Alstonia scholaris (L.) R. Br. "milky bean", "milky pine" [MSM, Julatten, 
Qld] 
*Tabernaemontana divaricata (L.) R. Br. "mock gardenia" [GS, Tolga, Qld] 
Ochrosia elliptica Labill. [GS, Tolga, Qld] _ 

Daphnis protrudens (Felder) 


RUBIACEAE 
Timonius timon (Spreng.) Merr. var timon [GS, Windsor Tableland, Qld] 


16 Australian Entomologist, 1998, 25 (1) 


Eupanacra splendens (Rothschild) 
Add to the records listed by Moulds (1984) the following: 


ARACEAE 
*Monstera deliciosa Liebm. "monstera" [Dennis Kitchen, JO, Cairns, Qld] 


Rhaphidophora australasica F. M. Bailey [H. Beste, Julatten, Qld] 


Hippotion boerhaviae (F.) 
RUBIACEAE 
Hedyotis sp. [JO, Trinity Beach, Qld] 


*Pentas lanceolata (Forssk.) Deflers "pentas" [JO, Trinity Beach, Qld - 
larvae were not found naturally on Pentas but readily accepted it when 
transferred from Hedyotis] 


Hippotion brennus (Stoll) 
DILLENIACEAE 
Hibbertia scandens (Willd.) Gilg "golden guinea vine 
Kuranda, Qld] 


RUBIACEAE 
*Pentas lanceolata (Forssk.) Deflers "pentas" [JO, Trinity Beach, Qld] 


Pogonolobus reticulatus F. Muell. [JO, Trinity Beach, Qld] 


mow 


snake vine" [JO, 


Hippotion rosetta (Swinhoe) 
RUBIACEAE 
*Pentas lanceolata (Forssk.) Deflers "pentas" [D. Lane, Atherton, Qld] 


*Richardia brasiliensis Gomes "Mexican clove" "white eye" [AJG, Yorkeys 
‘Knob, Qld] 

*Richardia scabra L. [AJG, Yorkeys Knob, Qld] 

Spermacoce exserta Benth. [Cliff Meyer, Darwin, NT] 


VITACEAE 
Cayratia clematidea (F. Muell.) Domin "wild grape" [AJG, Yorkeys Knob, 
Qld] 
‘Hippotion scrofa (Boisduval) 
Add to the records of Moulds (1981, 1984) the following: 
ASTERACEAE 


*Xanthium spinosum L. "Bathurst burr" [G. Brown, Cootamundra, NSW - 
identification uncertain; record requires confirmation] 


RUBIACEAE 
Hedyotis sp. [JO, Trinity Beach, Qld] 


Spermacoce exserta Benth. {Cliff Meyer, Darwin, NT] 


Australian Entomologist, 1998, 25 (1) 17 


Hopliocnema brachycera (Lower) 
MYOPORACEAE 
Eremophila willsii F. Muell. (MSM, Wallara Stn, NT) 


Eremophila exotrachys Kraenzlin (MSM, Wallara Stn, NT) 


CASUARINACEAE 

A data label attached to a specimen in the South Australian Museum states 
that it was reared from Casuarina. As larvae feed on Eremophila it is most 
unlikely that Casuarina is a food plant and the record is here disregarded. 


Hyles livornicoides (Lucas) 
Add to the records listed in Moulds (1981) the following: 


FABACEAE 
*?Medicago [sativa] L. "lucerne" [Lower (1897) - a doubtful record that 
requires confirmation] 


Leucomonia bethia (Kirby) 
VERBENACEAE 
Clerodendrum floribundum R. Br. "lJollybush" [GS, Walsh River, Mareeba 
district, Qld] 


Macroglossum alcedo (Boisduval) 
RUBIACEAE 
Hodgkinsonia frutescens C. T. White [GS, MSM. Tolga, Wongabel S. F. and 
Yungaburra, Qld] 


Macroglossum dohertyi (Rothschild) 
RUBIACEAE 
Myrmecodia platytyrea subsp. antoinii (Becc.) C. R. Huxley & Jebb "ant 
plant" [D. Lane, Iron Range, Qld] 


Macroglossum micaceum (Walker) 
RUBIACEAE 
Canthium sp. [GS, JO, Forty Mile Scrub, Qld] 


Macroglossum prometheus (Boisduval) 


RUBIACEAE 
Morinda citrifolia L. "cheesefruit" "great morinda" [AJG, Yorkeys Knob, 
Qld] 
Macroglossum tenebrosa (T.P. Lucas) 
RUBIACEAE 


Morinda salomonensis Engl. [D. Kitching, Kuranda, Qld] 


Macroglossum vacillans (Walker) 
LOGANIACEAE 


18 Australian Entomologist, 1998, 25 (1) 


Strychnos lucida R. Br. "strychnine tree" [GS, Myall Ck (=York Downs), 
Cape York Pen., Qld] 


Meganoton rufescens (Butler) 
Add to the records listed in Moulds (1984) the following: 


ANNONACEAE 
*Annona muricata L. "soursop" [R. Straatman and MSM, Kuranda, Qld] 


Psilogramma menephron menephron (Cramer) 
Add to the records listed in Moulds (1981, 1984) the following: 


BIGNONIACEAE 
Deplanchea tetraphylla (R. Br.) F. Muell. "golden bouquet tree" [JO, Trinity 
Beach, Qld] 


*Radermachera sinica (Hance) Hemsl. [J. McMaugh and C. Cassar, Sydney, 
NSW] 


CASUARINACEAE 

Walker (1856) formalized the name Macrosila casuarinae from a Boisduval 
manuscript name; this is now considered a junior synonym of Psilogramma 
menephron. Boisduval [1875] gives Casuarina as a larval food plant. A 
complete absence of other Australian records for this common moth from 
such an abundant plant suggests Boisduval's record is erroneous. Further, I 
have been unable to persuade young larvae to feed on Casuarina. However, 
Robinson.,(1975) records Psilogramma jordana B-Bkr in Fiji as feeding on 
Casuarina nodiflora. Despite this contradiction I believe the evidence 
dismissing Casuarina as a food plant for P. menephron outweighs the 
likelihood of this ancient record being correct and I here disregard the record 
in the absence of further evidence. 


OLEACEAE 
Jasminum didymum subsp. lineare (R. Br.) P. S. Green [MSM, Townsville 
district, Qld] 


*Osmanthus fragans (Thunb.) Lour. "fragrant olive" [J. McMaugh, Sydney, 
NSW] 


VERBENACEAE 
*Citharexylum hydalglense [A. B. Rose, Forster, NSW] 


Clerodendrum cunninghamii Benth. [GS, Mt Gravatt, Qld] 


Clerodendrum tomentosum (Vent.) R. Br. "lolly bush" [C. N. Smithers, 
Sydney, NSW] 


*Duranta repens L. "golden-dewdrop" "pigeon berry" [A. B. Rose, Forster; 
J. McMaugh, Sydney, NSW] 


Vitex glabrata R. Br. [Cliff Meyer, Kununurra, WA] 


Australian Entomologist, 1998, 25 (1) 19 


Vitex trifolia L. [Gordon Jones, Roebourne, WA; J. McMaugh, Sydney, 
NSW] 


Synoecha marmorata Rothschild & Jordan 
MYOPORACEAE 
Eremophila mitchellii Benth. "sandalbox", "budda", "false sandalwood", 
"bastard sandalwood", "emu bush" [Lucas 1891] 


Theretra latreillii (W.S. Macleay) 
Add to the records listed in Moulds (1981, 1984) the following: 


RUBIACEAE 

*Pentas lanceolata (Forssk.) Deflers "pentas" [JO, Cairns, Qld - larvae were 
not found on pentas but those transferred from other food plants readily 
accepted it.] 


VITACEAE 
Cissus opaca F. Muell. "pepper vine" [J. Moss, Brisbane, Qld] 


Theretra oldenlandiae (F.) 
Add to the records listed in Moulds (1981, 1984) the following: 


ARACEAE 
*Zantedeschia aethiopica (L.) K. Spreng. "arum lily" [MSM, Brisbane, Qld] 


RUBIACEAE 
Hedyotis sp. [JO, Trinity Beach, Qld] 


Theretra queenslandi (T.P. Lucas) 
URTICACEAE 
Dendrocnide excelsa (Wedd.) Chew "giant stinging tree" "fibrewood" [Clyne 
(1980); John Stockard, Wingham, NSW; AJG, Toowoomba, Qld; A. Hiller, 
Mt Glorious, Qld] 


Dendrocnide moroides (Wedd.) Chew "gimpi gimpi" "gympie" [A. Walford- 
Huggins, Tully Falls, Qld] 


Dendrocnide photinophylla (Kunth) Chew "shining-leaved stinging tree" 
"mulberry-leaved stinging tree" "fibrewood" [GS, Mt Tamborine, Qld; Anne 
Garrett, Rockhampton district, Qld.] 


Pipturus argenteus (G. Forst.) Wedd. "native mulberry" [GS, Tolga, Qld; 
AG, Kuranda, Qld.] 


Theretra silhetensis (Walker) 
ONAGRACEAE i 
*Lugwigia octovalvis (Jacq.) P. H. Raven "water primrose" [J. Moss, 
Brisbane, Qld; GS, Eurimbulah, Qld] 


RUBIACEAE 
Hedyotis sp. [JO, Trinity Beach, Qld] 


20 Australian Entomologist, 1998, 25 (1) 


*Pentas lanceolata (Forssk.) Deflers "pentas" [JO, Trinity Beach, Qld - 
larvae not found naturally on pentas but readily accepted it when transferred 
from Hedyotis] 


VITACEAE 
Cayratia clematidea (F. Muell.) Domin [AJG, Yorkeys Knob, Qld] 


Discussion 

Forty-three of the 64 Australian hawk moth species now have larval food- 
plant records of Australian origin. A further five species have food-plant 
records from localities beyond Australia, viz.: Hippotion rosetta (Swinhoe) 
on Boreria and Oldenlandia (Holloway 1987); Macroglossum corythus 
(Walker) on Strychnos, Guettarda, Morinda and Paederia (Holloway 1987); 
M. heliophila (Boisduval) on Morinda and Psychotria (Holloway 1987); M. 
insipida (Butler) on Hedyotis, Borreria, Spermacoce and Corchorus 
(Holloway 1987); and Theretra nessus (Drury) on Dioscorea, Ipomoea, 
Amaranthus, Impatiens, Citrullus, Arachis, Boerhavia, Knoxia, Morinda, 
Oldenlandia, Spermacoce, Glossostigma and Camellia (Holloway 1987, 
Mackey 1975, Seitz 1928-29). Food-plant records for the remaining 15 
Australian hawk moth species are lacking. 


Known Australian food plants now total 196 species in a remarkable 122 
genera and 43 families. The Rubiaceae (34 spp) has by far the highest 
representation followed by Bignoniaceae (15 spp), Vitaceae (15 spp) and 
Oleaceae (14 spp). The Rubiaceae support 22 Australian hawk moth species, 
more than for any other plant family. In comparison, the Vitaceae support 11 
species but the Bignoniaceae only four and the Oleaceae just one. 


One third of the known Australian food plants are introduced, 66 species in 
all spread across 23 families. Previously I have discussed the polyphagous 
nature of some Australian hawk moths (Moulds 1981), notably 
Gnathothlibus erotus (Cramer), Hippotion celerio (L.), Psilogramma 
menephron, Theretra latreillii and T. oldenlandiae. These five species 
account for 49 of the introduced plants and P. menephron alone for 25. 
These moths are all wide-ranging species with distributions extending at least 
throughout the Indo-Australian region. 


The 17 hawk moths endemic to Australia (a quarter of all Australian species) 
together feed on 66 species in 21 families but only nine of these plants are 
exotics. Coenotes eremophilae is an exception amongst the Australian 
endemic hawk moths, in that it has a large food-plant range (27 species in 13 
families) accounting for 26 of the 66 species, and 7 of the 21 families above. 
However, the plants that it feeds on are in families that are for the most part 
closely related, and all but three are Australian natives. 


To draw conclusions at this time about the diversity of Australian hawk moth 
food plants would be premature in view of the incomplete knowledge 
available. However, two trends are worth noting. Firstly, those hawk moths 


Australian Entomologist, 1998, 25 (1) 21 


with the widest distributions also tend to have the broadest range of food 
plants. Hippotion celerio, for example, is a cosmopolitan moth reaching the 
British Isles and North America, and from Australia alone it is recorded 
feeding on 25 species in 10 families. Among the Australian endemics 
Coenotes eremophilae has by far the widest distribution occurring throughout 
much of mainland Australia, and it has by far the highest food plant diversity 
as noted above. An exception to this trend seems, at first, to be Agrius 
convolvuli (L.) which, like H. celerio, is cosmopolitan but has comparatively 
few Australian food-plant records. However, throughout its world-wide 
distribution its larvae feed on a wide range of plants covering 7 families 
(Holloway 1987). In contrast, narrow-range species such as Hopliocnema 
brachycera, Synoecha marmorata (T. P. Lucas) and Coequosa triangularis 
are each known to feed on plants in just one family. 


Secondly, there is a tenuous correlation between the abundance of a species 
and its food-plant diversity. Common species tend to have a range of food 
plants spanning several families. For example, Psilogramma menephron (38 
species in 6 families), Theretra oldenlandiae (25 species in 8 families) and 
Gnathothlibus erotus (17 species in 6 families). Less abundant hawk moths 
have food plants which tend to be closely related. For example Coequosa 
triangularis is known to feed on nine different plants but all within the 
Proteaceae. The seven Australian Macroglossum species for which food 
plants are known are all uncommon; together they have 10 known food 
plants, nine of which are in the Rubiaceae and one in Loganiaceae. 


One would expect some kind of correlation between the systematic positions 
of hawk moths and their food plants but there is no obvious broad-based 
association in this regard. This subject is a complex one and falls beyond the 
scope of this paper and is not pursued here. 


Acknowledgments 

I am sincerely grateful to those who have provided food plant records: Hans 
Beste, Graham Brown, C. Cassar, Densey Clyne, Garry Fitt, Jim Frazier, 
Anne Garrett, Alan Graham, the late E. A. Henty, Tony Hiller, Gordon Jones, 
Dennis Kitchen, David Lane, N. Marks, Judy McMaugh, Cliff Meyer, John 
Olive, Clive Pratt, Tony Rose, Garry Sankowsky, John Stockard, Courtenay 
Smithers, the late Ray Straatman, Peter Valentine, Allan Walford-Huggins 
and Bruce White. The assistance of Tony Irvine and Garry Sankowsky in 
providing plant identifications is gratefully acknowledged. I am also 
indebted to my wife Barbara for typing the manuscript. 


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WALKER, F. 1856. List of the specimens of lepidopterous insects in the collection of the 
British Museum. Part 8, Sphingidae. British Museum, London. 271 pp. 


Australian Entomologist, 1998, 25 (1): 23-27 23 


PHACONEURA (HOMOPTERA: MEENOPLIDAE) 
ATTENDED BY ANTS OF THE GENUS PARATRECHINA 
(HYMENOPTERA: FORMICIDAE) IN CAVES 


W.F. HUMPHREYS 


Western Australian Museum, Francis Street, Perth, WA 6000 


Abstract 

Cave inhabiting species of Phaconeura Kirkaldy from the Kimberley and Cape Range, Western 
Australia, are attended by ants of the genus Paratrechina Motschoulsky, which themselves 
probably are especially adapted for cave life. 


Introduction 

The Australian tropics have been found recently to contain a diverse 
troglobitic fauna, in both humid (Howarth 1988) and arid (Humphreys 
1993b) regions. One of the more diverse groups of animals found in the 
caves is planthoppers (Homoptera) of the fulgoroid families Cixiidae 
(especially the genus Solonaima Kirkaldy: Hoch 1988, Hoch and Howarth 
1989a, 1989b, 1989c) and Meenoplidae (especially species of the genus 
Phaconeura Kirkaldy: Hoch 1990, 1993). By 1989 North Queensland had 
recorded the highest concentration of cave-adapted Fulgoroidea in the world 
(Hoch and Asche 1989). 


Until 1990 only one cave adapted meenoplid was known in Australia, 
Phaconeura pluto Fennah from Nambung National Park in Western Australia 
(Fennah 1973). Subsequently four new species of Phaconeura were 
described from North Queensland caves (Hoch 1990) and another (Hoch 
1993) from caves in the Cape Range of Western Australia (Humphreys 
1993a, 1993b). Since then several more undescribed cavernicolous species 
of Phaconeura (H. Hoch, pers. comm.) have been found in the area of Cape 
Range and in the Kimberley. Cave-adapted meenoplids have been found 
exclusively in limestone caves in Australia, whereas elsewhere they are only 
known from lava tubes (Hoch 1990) in the Canary Islands (Remane and 
Hoch 1988) and Western Samoa (Hoch and Asche 1988). Ant-Homoptera 
associations are well known (Schaefer 1987, Bourgoin in press) but have so 
far been reported only from epigean species. 


The distribution of obligatory cave dwelling (troglobitic) apterous species of 
Phaconeura, between isolated but neighbouring towers of the Chillagoe 
Karst, led Howarth (pers. comm. in Hoch 1990) to presume the species to be 
attended by ants in order to explain their dispersal; species of Paratrechina 
Motschoulsky occur in these caves (Howarth 1988). Here support is 
provided for Howarth's supposition by reporting the attendance of 
meenoplids by ants in caves in the Kimberley and Cape Range, Western 
Australia. 


24 Australian Entomologist, 1998, 25 (1) 


Observations 

During surveys of cave fauna in northwestern Australia (Humphreys 1993c, 
1995) meenoplids were found to occur widely but the observations reported 
here are from two main caves in the Kimberley and Cape Range. 


Cave KNI-9 is in the Ningbing Range, Kimberley (15°17'S; 128°37'E), 
formed in the exposed Devonian reef system. The cave is mainly horizontal 
and is ca 136 m long; it is one of many in the southern Ningbing Range. 


Cave C-29 is in Cape Range, Carnarvon Basin (22°06'S; 114°O1'E) in an 
anticline of Miocene limestones. The cave comprises a 24 m deep chamber 
from which two passages extend for 105 m (R. D. Brooks, pers. comm.) and 
where the ants and meenoplids were found. The chamber is often dry but 
seepages sometimes moisten the cave soil at the extremities of the passages. 


In the caves meenoplids are typically found wandering over the substrate or 
feeding on fine growing roots of undetermined plants, probably often Ficus, 
usually on damp soil covered surfaces. However, at two places in cave KNI- 
9, nymphs of a yet undescribed species of Phaconeura (H. Hoch, pers. 
comm. 1994) were found in groups on roots, either on the surface or in soil 
cavities, with ants in attendance which appeared to groom the nymphs. The 
ants became agitated when disturbed and began to collect the nymphs, both 
from the surface roots and the underground cells, and transport them away 
from the area. Within several minutes after disturbance no nymphs were left 
in the area. In Cape Range (C-29) nymphs of Phaconeura sp. were found in 
groups on roots, either on the surface or in soil cavities, with ants in 
attendance (A. Amarkey, pers. comm.); nymphs cannot be determined to 
species but Phaconeura proserpina Hoch is known from nearby caves. 


In both areas the ants were species of Paratrechina, a genus that requires full 
revision before species-level identification will be possible (S. Shattuck, pers. 
comm. 1994). These ants have very small eyes and it "seems likely that this 
is one of the few ants known that is especially adapted for cave life" (S. 
Shattuck, pers. comm. 1995). Small-eyed species of Paratrechina have also 
been collected from Cutta Cutta Cave, N.T., in the same parts of the cave as 
Phaconeura sp. indet. (W. Binks, R. D. Brooks and B. Vine, pers. comm.), 
although no clear association was reported. 


Paratrechina is widely distributed, although much more common in eastern 
and northern areas of Australia (S. Shattuck, pers comm. 1994). While 
Paratrechina have been considered to be opportunists (Reichel and Andersen 
1996), members of the genus are known to attend Hemiptera. The sugarcane 
mealybug Saccharicoccus sacchari (Cockerell) (Pseudococcidae), is 
consistently attended both above and below ground by ants, including 
Paratrechina sp. prob. vaga (Forel), which were observed carrying the 
mealybugs underground (Carver et al. 1987). In addition Paratrechina 
obscura Mayr was involved in behaviour - removing mummies from nodes - 
that has been interpreted as mutualistic (De Barro 1990). 


Australian Entomologist, 1998, 25 (1) 25 


Microclimate 

The meenoplids were found in those parts of the cave that contained root 
systems and where the air was nearly saturated with water (Table 1). In KNI- 
9 these areas were in high parts of the cave where the warmer air collected. 
In C-29 the meenoplids were found at the extremities of the tunnels where 
humidity is greatest. The fauna of tropical caves, even in arid zones, is 
dependent on high moisture levels (Howarth 1980, Humphreys and Collis 
1990, Humphreys 1991, Weinstein 1994). 


Table 1. The temperature (°C) and relative humidity (%) associated with cave KNI-9 
in May and June 1994. The mean is given with its standard deviation in parentheses. 
Temperature and relative humidity were spot measured using a whirling hygrometer 
(Brannan, England). 


Location n °C % 
Outside cave 3 29.2 (1.72) 44 (16) 
Inside cave: no meenoplids 26 25.9 (1.76) 86 (15.9) 

meenoplids 3 27.3 (0.46) 98 (0.6) 
Discussion 


The fauna found in the Kimberley caves is not predominantly troglobitic, 
unlike that of tropical caves elsewhere in Australia (Howarth 1988, 
Humphreys 1993b, 1993c). Only one of 14 species (7%) associated with the 
meenoplids was troglomorphic, that is it had morphological adaptations 
associate with cave life, namely Blattodea [Nocticola brooksi Roth 
(Blattodea: Nocticolidae)]. By contrast, arid Cape Range contains a very rich 
cave fauna with rainforest affinities (Humphreys 1993b, 1993c) and, of the 
species known from C-29, half (n=8) were highly troglomorphic, namely 
Nocticola flabella Roth (Blattodea: Nocticolidae), Ngamarlanguia luisae 
Rentz & Su (Orthoptera: Gryllidae: Nemobiinae), Stygiochiropus communis 
Humphreys & Shear (Diplopoda: Paradoxosomatidae) and Draculoides vinei 
(Harvey) (Arachnida: Schizomida). 


Cave restricted animals ultimately feed on photosynthetically derived energy, 
except in special cases of chemoautotrophy. The energy is transferred to the 
cave largely by water, as fine or gross plant material, by animals, such as bats 
and rhaphidophorid cave crickets foraging outside the cave, and by plants by 
means of root growth and sap transport. Cave restricted animals may eat the 
roots alive or dead, feed on root exudates or directly on the sap, as do 
meenoplids. The presence in these caves of a two level assemblage of 
terrestrial species utilizing plant roots may be the first step to recognising 
more complex root feeding assemblages in caves, as found recently for 


26 Australian Entomologist, 1998, 25 (1) 


aquatic invertebrates for which rich assemblages are supported by root mats 
in caves (Jasinska et al. 1996). 


Acknowledgments 

It is my pleasure to acknowledge the identifications and comments of 
Hannelore Hoch and Steve Shattuck and the invaluable support of Julianne 
Waldock, Darren Brooks and Brian Vine. Facilities were made available by 
Bob Shackles, Officer in Charge, Frank Wise Institute for Tropical 
Agricultural Research in Kununurra. Professor Dr Hannelore Hoch and Dr 
M. B. Malipatil identified Emesinae. Field work was funded under the 
National Estate Grants Scheme. 


References 
BOURGOIN, T., in press. Habitat and ant attendance in Hemiptera Auchenorrhyncha, a 
phylogenetic test. Memoires du Museum National d'Histoire Naturelle (Zoologie). 


CARVER, M., INKERMAN, P.A. and ASHBOLT, N.J. 1987. Anagyrus saccharicola 
Timberlake (Hymenoptera: Encyrtidae) and other biota associated with Saccharicoccus 
sacchari (Cockerell) (Homoptera: Pseudococcidae) in Australia. Journal of the Australian 
Entomological Society 26: 367-368. 


DE BARRO, P.J. 1990. Natural enemies and other species associated with Saccharicoccus 
sacchari (Cockerell) (Hemiptera: Pseudococcidae) in the Bundaberg area, southeast 
Queensland. Journal of the Australian Entomological Society 29: 87-88. 


FENNAH, R.G. 1973. ` Three new cavernicolous species of Fulgoroidea (Homoptera) from 
Mexico and Western Australia. Proceedings of the Biological Society, Washington 86: 439- 
446. 


HOCH, H. 1988. Five new epigean species of the Australian planthopper genus Solonaima 
Kirkaldy (Homoptera: Fulgoroidea: Cixiidae). The Beagle, Records of the Northern Territory 
Museum of Arts and Science 5: 125-133. 

HOCH, H. 1990. Cavernicolous Meenoplidae of the genus Phaconeura (Homoptera: 
Fulgoroidea) from Australia. Occasional Papers of the Bishop Museum 30: 188-203. 

HOCH, H. 1993. A new troglobitic planthopper species (Homoptera: Fulgoroidea: 


Meenoplidae) from Western Australia. Records of the Western Australian Museum 16: 393- 
398. 


HOCH, H. and ASCHE, M. 1988. A new troglobitic meenoplid from a lava tube in Western 
Samoa (Homoptera: Fulgoroidea: Meenoplidae). Journal of Natural History 22: 1489-1494. 
HOCH, H. and ASCHE, M. 1989. Cave-dwelling planthoppers of Australia (Insecta: 
Homoptera: Fulgoroidea). Jn Pearson, L. (ed.). Tropicon Conference, Lake Tinaroo, Far North 
Queensland. 27-31 Dec. 1988. 67-75. Australian Speleological Federation, Cairns. 

HOCH, H and HOWARTH, F.G. 1989a. Six new cavernicolous cixiid planthoppers in the 
genus Solonaima from Australia (Homoptera: Fulgoroidea). Systematic Entomology 14: 377- 
402. 

HOCH, H. and HOWARTH, F.G. 1989b. Reductive evolutionary trends in two new 
cavernicolous species of a new Australian cixiid genus (Homoptera: Fulgoroidea). Systematic 
Entomology 14: 179-196. 


Australian Entomologist, 1998, 25 (1) 27 


HOCH, H. and HOWARTH, F.G. 1989c. The evolution of cave-adapted cixiid planthoppers in 
volcanic and limestone caves in North Queensland, Australia (Homoptera: Fulgoroidea). 
Mémoires de Biospéologie 16:17-24. 

HOWARTH, F.G. 1980. The zoogeography of specialised cave animals: a bioclimatic model. 
Evolution 34: 394-406. 

HOWARTH, F.G. 1988. Environmental ecology of north Queensland caves: or why there are 
so many troglobites in Australia. Jn Pearson, L. (ed.) 17th biennial conference, Australian 
Speleological Federation Tropicon Conference, Lake Tinaroo, Far North Queensland 27-31 
Dec. 1988. 76-84. Australian Speleological Federation, Cairns. 

HUMPHREYS, W.F. 1991. Experimental re-establishment of pulse-driven populations in a 
terrestrial troglobite community. Journal of Animal Ecology 60: 609-623. 

HUMPHREYS, W.F. 1993a. The significance of the subterranean fauna in biogeographical 
reconstruction: examples from Cape Range peninsula, Western Australia. Records of the 
Western Australian Museum, Supplement 45: 165-192. 


HUMPHREYS, W.F. 1993b. Cave fauna in semi-arid tropical Western Australia: a diverse 
relict wet-forest litter fauna. Mémoires de Biospéologie 20: 105-110. 

HUMPHREYS, W.F. (ed.). 1993c. The biogeography of Cape Range, Western Australia. 
Records of the Western Australian Museum, Supplement 45: 1-248. 

HUMPHREYS, W.F. 1995. Limestone of the east Kimberley, Western Australia - karst and 


cave fauna. Report to the Australian Heritage Commission and the Western Australian 
Heritage Committee. 190 pp.+ xix. Unpublished. 


HUMPHREYS, W.F. and COLLIS, G. 1990. Water loss and respiration of cave arthropods 
from Cape Range, Western Australia. Comparative Biochemistry and Physiology 95A: 101- 
107. 


JASINSKA, E.J., KNOTT, B. and McCOMB, A.J. 1996. Root mats in ground water: a fauna- 
rich habitat. Journal of the North American Benthological Society 15: 508-519. 

REICHEL, H. and ANDERSEN, A.N. 1996. The rainforest ant fauna of Australia's Northern 
Territory. Australian Journal of Zoology 44: 81-95. 

REMANE, R. and HOCH, H. 1988. Cave-dwelling Fulgoroidea (Homoptera 
Auchenorrhyncha) from the Canary Islands. Journal of Natural History 22: 403-412. 
SCHAEFER, C.W. 1987. The early habitat of the Auchenorrhyncha. In C. Vidano and A. 


Arzone (eds). Proceedings of the 6th Auchenorrhyncha Meeting, Turin, Italy, Sept. 1987, 135- 
146. 


WEINSTEIN, P. 1994. Behavioural ecology of tropical cave cockroaches: preliminary field 
studies with evolutionary implications. Journal of the Australian Entomological Society 33: 
367-370. 


28 


Australian Entomologist, 1998, 25 (1) 


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Australian Entomologist, 1998, 25 (1): 29-31 29 


NOTES ON THE DISTRIBUTION OF THE DINOSAUR ANT 
NOTHOMYRMECIA MACROPS CLARK (HYMENOPTERA: 
FORMICIDAE) IN SOUTH AUSTRALIA 


C.H.S. WATTS, A.J. McARTHUR and R. FOSTER 
South Australian Museum, North Terrace, Adelaide, SA 5000 
Abstract 


Nothomyrmecia macrops Clark, the only surviving member of the ant subfamily 
Nothomyrmeciinae and known previously from one restricted locality in South Australia and 
one site in Western Australia, is recorded from 17 additional localities over a linear distance of 
more than 400 km in northern Eyre Peninsula, South Australia. It appears to be a relatively 
common nocturnal species, active only on relatively cold nights in regions of mature mallee. 
Introduction 

Nothomyrmecia macrops Clark is the only surviving member of the ant 
subfamily Nothomyrmeciinae. It retains a large number of primitive 
morphological and behavioural features (Holldobler and Taylor 1983) and 
hence has attracted the common name ‘dinosaur ant’ or ‘living fossil ant’. 


First recorded in 1934 from the vicinity of Esperance in Western Australia, it 
was not collected again until it was discovered at Poochera on Eyre 
Peninsula, South Australia in 1976 (Taylor 1978). Since then searches by 
Taylor and others have failed to locate additional colonies (R. W. Taylor, 
pers. comm.). The future of this phylogenetically isolated and hence 
scientifically important ant was in doubt. 


Methods 

During the spring and early summer of 1995, with some guidance from Dr R. 
W. Taylor, we conducted a search for additional colonies of N. macrops in 
the Upper Eyre Peninsula region of South Australia (Fig. 1). 


Trials of various survey methods (beating, hand searching, the use of 
‘tanglefoot™’, pitfall traps and baiting with honey) were undertaken on the 
known populations at Poochera. All of the above methods produced N. 
macrops except the use of pitfalls. Beating bushes and baiting tree trunks 
with honey proved equally effective. However, in terms of unit effort, 
baiting was clearly superior, being a rapid and easy technique capable of 
quickly surveying relatively large areas. 


It may be indicative of the ant’s behaviour that no specimens were caught in 
pitfall traps, although the traps were exposed for two months and were 
scattered among the mallee, often only a metre or two from trees on which 
the ant was observed climbing on most nights. 


As previously demonstrated by Hölldobler and Taylor (1983), N. macrops is 
nocturnal and is generally active only at temperatures below about 15°C 
(although two individuals were caught on different nights at 20°C). 


30 Australian Entomologist, 1998, 25 (1) 


Based on our experiences at Poochera we adopted the following survey 
method. During the day we located likely patches of mallee scrub. In the 
late afternoon up to five sites, usually several kilometres apart, were baited 
by smearing honey bait on tree trunks around eye height whilst walking in a 
straight line or loop for twenty minutes, resulting in a bait trail of 200-500 m 
on 50-80 tree stems. Soon after dusk the bait line was visited and samples of 
any ants present collected. In general, sites were surveyed on only one night. 








130 135 


Fig. 1. Eyre peninsula (South Australia) survey sites for Nothomyrmecia macrops. 
Filled squares indicate sites where N. macrops was collected. Shaded area refers to 
open scrub woodland as given in Griffin and McCaskill 1986. (Because of the scale 
of the map not all sites are distinguished). 


Results 

Seventy-four separate sites were surveyed between Lake Gilles in the east 
and Nundroo in the west. Nothomyrmecia macrops was found at 17 sites in 
addition to the known sites at Poochera (Fig. 1). (More specific details of 
localities of sites surveyed are available from A. J. McArthur). Negative 
results do not, of course, necessarily imply that it was not present. This is 
particularly true for a species such as N. macrops which is known to be 
inactive on warm nights and to be erratic in activity even at low 
temperatures. 


Australian Entomologist, 1998, 25 (1) 31 


We did not attempt to measure the density of N. macrops, nor the geographic 
extent of the individual colonies we located. Our impression was that where 
it occurred it was in reasonable numbers, with 10-12 individuals at a bait 
station not unusual. 


To the best of our ability in the time available, we attempted to describe in 
general terms each of the sites visited. Nothomyrmecia macrops appeared to 
be associated with sites characterised by the following: ‘old growth’ mallee, 
long unburnt with a mixture of tree sizes and at least a few large old trees; 
mallee dominated by Eucalyptus oleosa, E. brachycalyx and E. gracilis alone 
or more often in combination; loose, friable calcareous soils with a high 
‘fines’ content; fairly bare ground with a thin, flat layer of litter and little 
understorey and a high diversity of ant species, but no dominant aggressive 
species. 


Discussion 

We found N. macrops to be easily surveyed using a dilute honey bait on cool 
(<18°C) nights. Using this technique we found N. macrops to be a relatively 
common species in areas of ‘old growth’ patchy mallee with sparse 
understorey and a thin litter layer on friable soil, between Lake Gilles and 
Penong on Eyre Peninsula, a distance of about 400 km. Within this area it 
occurred as a member of several ant assemblages. We expect that the 
distribution of the species will eventually be shown to be even greater. We 
see no reason to consider the species endangered as long as no further major 
clearing of mallee vegetation occurs. Remnant roadside mallee represents a 
significant proportion of its remaining habitat. The species occurs in the 
Lake Gilles Conservation Park and the Chadinga Conservation Reserve. 


Acknowledgments 

This work was supported by a grant from the Endangered Species Unit of 
Environment Australia. Dr R. W. Taylor provided valuable advice from his 
deep knowledge of the behaviour of this ant. Mr Adrian (Bill) Pyke is 
thanked for his considerable contribution to the field work. Plant 
identifications were done by the South Australian Herbarium. Collected 
material is lodged in the South Australian Museum. 


References 
GRIFFIN, T. and McCASKILL, M. 1986. Atlas of South Australia. Government Printer, 
Adelaide. 


HOLLDOBLER, B. and TAYLOR, R.W. 1983. A behavioural study of the primitive ant 
Nothomyrmecia macrops Clark. Insectes Sociaux 30: 384-401. 


TAYLOR, R.W. 1978. Nothomyrmecia macrops: a living-fossil ant rediscovered. Science 
201: 979-984. 


32 Australian Entomologist, 1998, 25 (1) 


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FOR SALE: Butterflies from all parts of the world. Papua New Guinea, 
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Papilionidae inc. Parnassius, Delias, Charaxes etc. Free catalogue. David 
Hall, 6 Rule St, Cambridge Park, N.S.W., 2747. Ph. (047) 312 410. 


FOR SALE. 10-drawer wooden insect cabinets with recessed outer door and 
napthalene well. 560 x 400 x 36 mm. $200. G. Daniels 07 3365 
7084. E-mail: g.daniels @ento.uq.edu.au 


ENTOMOLOGICAL BOOKS. Pemberley Books are specialist suppliers of 
entomological literature across the world. Send for our free catalogue 
which lists a wide range of antiquarian, second-hand and new natural 
history titles. Pemberley Books, Ian Johnson, 34 Melrose Close, Hayes, 
Middlesex, UB4 OAZ, England. Tel/Fax: +44 181 561 5494. E-mail: 
ij @pembooks.demon.co.uk 


NOTES FOR AUTHORS 


Manuscripts submitted for publication should, preferably, be type-written, double spaced 
and in triplicate. Refer to recent issues for layout and style. 

All papers will be forwarded to two referees and the editor reserves the right to reject any 
paper considered unsuitable. 

Papers longer than eight printed journal pages will normally not be accepted. 

Papers will be accepted only if a minimum of 100 reprints is purchased. Manuscripts 
occupying less than one printed page may be accepted without charge if no reprints are 
required. Charges are as follows: cost per printed page $25 for 100 copies. Page charges 
may be reduced at the discretion of the Publications Committee. 

Illustrations: Black and white photographs must be submitted at the size they are to appear 
in the journal. Line drawings should be about twice their required size. 

Address papers to: The Editor 

The Australian Entomologist 
P.O. Box 537, 
Indooroopilly, Queensland, 4068 


Printed by Hans Quality Print, 20 Lyons Terrace, Windsor, Qld, 4030 


THE AUSTRALIAN 


Entomologist 


Volume 25, Part 1, 5 June1998 
KEX 
CONTENTS 


EASTWOOD, R. and KING, AJ. 
Observations on the biology of Arhopala wildei Miskin (Lepidoptera: Lycaenidae) 
-and its host ant Polyrhachis queenslandica Emery (Hymenoptera: Formicidae) 


WILLIAMS A.ALE., WILLIAMS, M.R. and HAY, R.W. 
A new species of Trapezites Hübner (Lepidoptera: Hesperiidae) from 
Western Australia - 


MOULDS, M.S. 
New larval food plants for Australian hawk moths (Lepidoptera: Sphingidae) 


HUMPHREYS, W.F. 
Phaconeura (Homoptera: Meenoplidae) attended by ants of the genus 
Paratrechina (Hymenoptera: Formicidae) in caves 


WATTS, C.H.S. , McARTHUR, AJ. and FOSTER, R. 


Notes on the distribution of the dinosaur ant Nothomyrmecia macrops Clark 
(Hymenoptera: Formicidae) in South Australia 


CONFERENCE NOTICES 
Australian Entomological Society Conference and Australasian Applied 
Entomological Research Conference 


RECENT LITERATURE 
An accumulative bibliography of Australian entomology 


_ ENTOMOLOGICAL NOTICES Inside back cover. 


ISSN 1320 6133 


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